Vegetative and reproductive morphology of Othniophyton elongatum (MacGinitie) gen. et comb. nov., an extinct angiosperm of possible caryophyllalean affinity from the Eocene of Colorado and Utah, USA

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Abstract
Eocene foliage formerly attributed to the extant araliaceous genus Oreopanax was found attached to twigs bearing inflorescences and infructescences unlike those of Araliaceae. Using newly observable characters of phyllotaxy, vegetative and floral buds, infructescences and seeds, we sought to reassess the affinities of this strange angiosperm. Fossils were studied from the Parachute Creek Member of the Green River Formation from near Bonanza, Utah and Douglas Pass, Colorado (ca. 47 Ma). Macrofossil impression remains were investigated by low angle reflected light and subtle details of the vegetative and floral buds, stamens, mature fruits, and seeds were revealed by optical shadow effect microscopy documenting previously obscure topographic surface features. Othniophyton elongatum (MacGinitie) Manchester, Judd, Correa-Narvaez gen. et comb. nov. has simple, short-petiolate, elongate, entire-margined leaves with thick midveins, pinnate, brochidodromous secondaries, common intersecondary veins and finely reticulate higher order venation. Inflorescences are small axillary cymes; flower buds are subglobose and pedicellate, in bract axils. The flowers are actinomorphic, bisexual, with ca. five imbricate perianth parts, ca. 24 stamens with elongate anthers and short filaments, arising from a hypanthium. The whorl of stamens persists to fruiting stage. The ovary is superior, with probable basal placentation and five stigmatic arms. Fruits are pedicellate berries with a cup-shaped hypanthium and contain ca. 15 lensoidal reniform seeds, each with a curved embryo and ornamented with concentric ridges. The combined characters refute the prior placement in Araliaceae, and rule out affinities with most extant clades of angiosperms. The distinctive combination of observed features does not coincide with any extant family. Among extant orders of Eudicots, the fossil seems to conform most closely to the order Caryophyllales, but key differences remain. This example indicates that the vegetation of ca. 47 million years ago included some taxa that cannot readily be placed in modern families and genera

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